Three New Records of Plant Parasitic Phyllosphere Fungi from Panama: Annellophora Phoenicis, Cercospora Corniculatae, and Sclerotium Coffeicola

14 1 93

NOTES ON GEOGRAPHIC DISTRIBUTION Check List 14 (1): 93–100 https://doi.org/10.15560/14.1.93

Three new records of plant parasitic phyllosphere fungi from Panama: Annellophora phoenicis, Cercospora corniculatae, and Sclerotium coffeicola

Roland Kirschner1, Orlando Cáceres2, Meike Piepenbring3

1 Department of Biomedical Sciences and Engineering, National Central University, Zhongda Rd. 300, Zhongli District, Taoyuan City 32001, Taiwan (R.O.C.). 2 Centro de Investigaciones Micologicas (CIMi), Departamento de Biologia, Universidad Autonoma de Chiriqui, David, Panama. 3 Department of Mycology, Goethe University Frankfurt am Main, Biologicum, Max-von-Laue-Str. 13, 60438 Frankfurt am Main, Germany. Corresponding author: Meike Piepenbring, [email protected]

Abstract Three fungi associated with living leaves of plants are new records for Panama: Annellophora phoenicis causing leaf spots of Cocos nucifera (Arecaceae), Cercospora corniculatae (C. apii s. lat.) on living leaves of Oxalis barrelieri (Oxalidaceae) with and without discoloration, and Sclerotium coffeicola on zonate leaf spots of Annona montana (An- nonaceae) and Dioscorea alata (Dioscoreaceae). Some records of A. phoenicis and S. coffeicola relevant for known geographical distribution and available by literature are critically revised.

Key words Anamorphic fungi; asexual fungi; Ascomycota; Basidiomycota; tropical fungi.

Academic editor: Roger Fagner Ribeiro Melo | Received 26 September 2017 | Accepted 11 December 2017 | Published 12 January 2018

Citation: Kirschner R, Cáceres O, Piepenbring M (2018) Three new records of plant parasitic phyllosphere fungi from Panama: Annellophora phoenicis, Cercospora corniculatae, and Sclerotium coffeicola. Check List 14 (1): 93–100. https://doi.org/10.15560/14.1.93

Introduction Methods Fruit production plays an important role in the economy Specimens were collected during July 2016 in Chiriquí of Panama. Bananas and coffee belong to the most Province, at the border of a small village in the lowland important products for export. Although plant protection of western Panama, approx. 150 m above sea level agencies need well-documented data about plant patho- (a.s.l.), with authorization from the Ministerio de Ambi- genic fungi, scientifically presented information about ente (MiAmbiente, Panama). Colors of symptoms on pathogens of cultivated plants is far from being complete leaves were observed using fresh material. Specimens of for Panama. Inventories of plant parasitic fungi, therefore, Sclerotium coffeicola were loaned from the Fungarium even recently, reveal new records for the economically Collections of the Herbarium of Royal Botanic Gardens, important fruit plants, such as Asterinella puiggarii on Kew (K). Fungal material was mounted in 10% KOH, guava (Piepenbring et al. 2011) and Ramichloridium with and without previous staining with aqueous Kongo biverticillatum and R. musae on banana (Kirschner and red solution, or in cotton blue in lactic acid and polyvinyl Piepenbring 2014). alcohol. Statistical treatments of the measurements are

Copyright Kirschner et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unre- stricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 94 Check List 14 (1) given with extreme values in brackets and mean value Notes. The species is identified based on the almost ± 1 standard deviation of n measurements. Specimens identical morphology described by Ellis (1958) and its were air-dried on an electrical dryer and deposited at occurrence on palms. This fungus has been recorded the Herbario de la Universidad Autónoma de Chiriquí, from Cocos and Phoenix species in the Old World (Ellis Panama (UCH). A sequence (603 bp) of the internal tran- 1971). In contrast to Ellis (1958, 1971), we rarely found scribed spacer (ITS) of the ribosomal RNA gene (rDNA) percurrent extensions of the conidium beak and we from a dried specimen of S. coffeicola was generated as observed inconspicuous, intercalary appressoria among described in Kirschner (2016), submitted to a BLAST superficial hyphal cells. A more detailed study, however, search and deposited in GenBank (MF170962). is required to reveal the exact mode of penetration into the leaf. In our opinion, a report as causative of palm leaf Results spot from Texas by Vann and Taber (1985), frequently cited in American textbooks (Horst 2001, Elliot et al. Ascomycota, incertae sedis 2004, Broschat et al. 2014), is based on a wrongly identified specimen, because the distinct sporodochia formed Annellophora phoenicis M.B. Ellis, Mycol. Pap. 70: 87 by stout conidiophores and the conspicuously verrucose (1958) conidia shown in the scanning electron microscopic pho- Figure 1 tograph, as well as gradual tapering of conidia from the Leaf spots amphigenous along the leaf margins, reddish base to the end instead of having an abrupt beak shown brown, with bright yellow margin, 10–35 × 3–7 mm, in the light microscopic photograph, clearly indicate that becoming confluent. Stromata lacking. Hyphae penetrat- the agent of palm leaf spot from Texas should be called ing into and through cell walls of the epidermis, forming Scolecostigmina palmivora (Sacc.) Kamal. Ellis (1958) small empty lacunae and cyanophilous areas within the described the type specimen of A. phoenicis from dead outer epidermis cell wall, intracellular hyphae incon- palm leaves so that it is not clear whether the fungus spicuous and rare in the epidermis, conspicuous in the was saprobic or parasitic. Reports from dead wood of mesophyll cells, pale brown, smooth, 1–3 µm wide. At Michelia baillonii (Magnoliaceae) in Thailand (Kodsueb the colony margin, hyphae are yellowish pale brown, et al. 2008) as well as from leaf litter of Anacardium smooth, occasionally verruculose, 1–2 µm wide, loosely occidentale (Anacardiaceae) in India (Shanthi & Vittal ramified and without appressoria; in the older part, the 2012) without any data on morphology may refer to other hyphae are densely ramified with numerous swollen inter- species. The considerably larger sizes of A. phoenicis var. calary appressoria showing a central round pale spot, 4–7 cubensis Hol.-Jech. described from a dead fallen branch × 2.5–4 µm. Conidiophores predominantly epiphyllous, of an undetermined tree in Cuba (Holubová-Jechová arising solitarily from a lobed foot cell (lobes only vis- 1988) would justify raising the variety to species level. ible in young conidiophores), being 10–15 wide and 4–5 The present concept of Annellophora, however, of being µm high, erect, straight, unbranched, cylindrical, brown, Sporidesmium-like but forming secondary conidia from smooth, two- to several-celled, with some intercalary percurrently extending conidial apices (Ellis 1958, 1971), cells darker than the other cells, length including foot cell appears artificial, because it comprises species from (10–)25–50(–65), width 4(–5) µm (n = 30). Conidiog- living leaves, dead wood, as well as other fungi (Ellis enous cells terminal, with percurrent extensions, truncate 1971). Since more recent reliable records of A. phoenicis at the apex, 4–11.5(–15) × 3–4 µm (n = 30). Conidia after Ellis (1971) and molecular sequence data are not broadly clavate or obclavate, sometimes with 1–2 con- available, detailed microscopic documentation of similar strictions, yellowish pale brown, smooth, distoseptate fungi associated with palm leaf spots is necessary in order in the main body, rarely forming percurrent extensions to approach correct species characterization. of the transversally 0–12-septate beak or a secondary conidium at the end of the beak, septa per conidium Ascomycota, Capnodiales altogether (9–)10–14(–16) (n = 30), distances between Cercospora corniculatae Hansf., Proc. Linn. Soc. septa 3–6 µm, size including beak (45–)56–75(–86) × London 155: 56 (1943) [1942–43], (= C. apii s. lat., (10–)10.5–12.5(–13) µm (n = 30), hilum 3–4 µm wide. Crous and Braun 2003) Specimen examined/new record. Panama, Chiriquí prov- Figure 3 ince, Corr. Dolega, Los Algarrobos, Casa de la Alemana, Leaf spots absent, sporulation taking place on green or garden, approx. 150 m a.s.l. (08°29′45″ N, 082°25′58″ diffusely reddish or yellow discolored leaves. Stromata W), on living leaves of Cocos nucifera L. (Arecaceae/ absent or reduced to few cells. Hyphae internal, pale Palmae), 23 July 2016, R. Kirschner 4292 (UCH). brown to hyaline, smooth, 2–4 µm wide. Conidiophores Known distribution (Fig. 2). “Malaya” = West Malaysia amphigenous, penetrating through stomata, solitary or in (Malaysia), Myanmar, New Guinea (Indonesia/Papua fascicles of up to 15, unbranched, rarely with one branch- New Guinea), Sierra Leone (Ellis 1971, Thaung 2008), ing at the base, cylindrical, geniculate, brown, smooth, India? (Shanthi and Vittal 2012), Thailand? (Kodsueb et 2–7-septate, septa 5–25 µm apart, (65–)86–157(–205) al. 2008), Panama (new record). × (5–)5.5–6.5(–7.5) µm (n = 30). Conidiogenous cells Kirschner et al. | Plant parasitic fungi from Panama 95

Figure 1. Annellophora phoenicis on Cocos nucifera. A. Leaf spots. B. Hypha that penetrated from appressorium into epidermis wall and cell. Lacuna in cell wall indicated with black arrow, cyanophilous cell wall reaction indicated with gray arrow, matrix with black arrowhead. C. External hyphae from colony margin with conidiophore bases (arrowheads). D. External hyphae from older part of colony with circles indicating penetration sites. E. Conidiophores. F. Conidia. Percurrent extension of two conidia is indicated by arrows. The three most right ones apically develop a secondary conidium. G. Detail of a conidium showing distosepta, pigmentation not shown. Scale bars: A = 1.5 cm, B = 5 µm, C–G = 10 µm. 96 Check List 14 (1)

intercalary and terminal, straight or geniculate, pale brown to brown, smooth, terminal ones (22–)31–50(–56) × 4–6(–7) µm (n = 30). Conidiogenous loci conspicuous, darkened, 1 µm thick, 2–3 µm wide. Conidia solitary, obclavate-cylindrical, straight or slightly undulate, sometimes the distal part curved, hyaline, smooth, 2–11-septate, (21–)51–100(–130) × 4–5 µm (n = 30), basal hilum thickened, darkened, 2–3 µm wide. Specimen examined/new record: Panama, Chiriquí province, Corr. Dolega, Los Algarrobos, approx. 150 m Figure 2. Annellophora phoenicis. World distribution, countries a.s.l. (08°29′45″ N, 082°25′58″ W), on living leaves reliably reported in the literature marked with green, Panama (new record) with brown red. Blue color indicates records considered of Oxalis barrelieri L. (Oxalidaceae), 25 July 2016, R. erroneous or doubtful in this study. Kirschner 4293 (UCH).

Figure 3. Cercospora corniculatae (C. apii s. lat.) on Oxalis barrelieri. A. Upper leaf surface covered by brown fascicles of conidiophores. B. Upper and lower leaf surfaces covered by brown conidiophores fascicles and white conidial mass. C. Fascicle of four conidiophores. D. Two of the rare basally branched conidiophores. E. Conidia. Scale bars = 20 µm. Kirschner et al. | Plant parasitic fungi from Panama 97

sporulation took place on non-discolored green as well as diffusely discolored leaves. The type of the invalid C. oxalidiphila Chupp & A.S. Mull. was studied by Braun (2000), who indicated the synonymy with C. corniculatae and suggested a close relationship or identity with C. apii. Species delimitation in the Cercospora apii species complex (Crous and Braun 2003) is still under progress (Groenewald et al. 2013).

Basidiomycota, Atheliales Sclerotium coffeicola Stahel, Bull. Dep. Landbouw Figure 4. Cercospora corniculatae (C. apii s. lat.). World distribution, countries reported in the literature marked with green, Panama Suriname 42: 15 (1921) (new record) with brown red. Figures 5, 6 Leaf spots zonate by alternating concentric pale and Known distribution (Fig. 4). Argentina, Myanmar, dark brown rings, with dark brown margin, 8–95 × 8–55 Puerto Rico, Trinidad and Tobago, Venezuela, Uganda, mm. Propagules hypophyllous, white, erect, lanceolate, USA (WI) (Crous and Braun 2003, Farr and Rossman smooth, 2–4 × 0.1–0.2 mm, easily detaching from the 2017), Panama (new record). leaf. Propagules composed of textura intricata at the base, over the length composed of parallel; scarcely branched, Notes. This is the first record of a Cercospora species hyaline, smooth or verruculose hyphae of 4–6 µm diam- on members of Oxalis in Panama. The other known eter; septa 20–70 µm apart, converging at the tip. Cercospora species on members of Oxalis, C. oxalidis A.S. Mull. & Chupp ex U. Braun & Crous, differs by its Specimen examined/new record. Panama, David, Corr. overall smaller dimensions from C. corniculatae/C. apii Dolega, Los Algarrobos, Casa de la Alemana, garden, (Crous and Braun 2003). Cercospora corniculatae has approx. 150 m a.s.l. (08°29′45″ N, 082°25′58″ W), on been recorded from O. barrelieri in Trinidad and Tobago living leaves of Annona montana Macfad. (Annonaceae), and Venezuela (Farr and Rossman 2017). On other 25 July 2016, R. Kirschner 4290 (UCH; ITS sequence: Oxalis species, the fungus was also recorded from fur- GenBank MF170962). ther countries (Crous and Braun 2003). In contrast to the Additional specimens examined. Sierra Leone, Njala descriptions of Cercospora spp. on Oxalis spp. (Chupp (Kori), on leaves of Jasminum multiflorum (Burm. f.) 1954, To-Anun et al. 2011), distinct leaf spots were not Andrews [as J. pubescens (Retz.) Willd.], 1933, F.C. present in the Panamanian specimen, but conspicuous Deighton M468, IMI 35109 (K, only sclerotia); same

Figure 5. Sclerotium coffeicola from Panama, fresh material on Annona montana, except D: on Dioscorea alata. A. Concentric zonate spots seen on the upper leaf side. B. Leaf spot seen from below showing the white lanceolate propagules. C. Lower leaf side showing propagules and orange gall midge larvae (blue arrows). D. Abaxial leaf spots of Dioscorea alata. E. Habitus sketch of propagules. F. Detail from the textura intricata of the base of the propagule. G. Detail from the parallel hyphae composing the main body of the propagule. H. Apex of propagule showing convergent hyphae. Scale bars: B = 1 cm, E = 1 mm, I = 100 µm, F, G = 20 µm. 98 Check List 14 (1)

Figure 6. Sclerotium coffeicola from Africa (Sierra Leone), collections from 1949 (K). A. Concentric leaf spot on the upper side of leaf of Aristolochia inflata (IMI 37953), a single propagule marked with arrow. B. Propagules from lower side of leaf of A. inflata (IMI 37953). C. Propagules from abaxial leaf side of Jasminum multiflorum (IMI 37954). Scale bars = 4 mm. area, on zonate leaf spots of Aristolochia inflata Kunth (as A. gibbosa Duch.), August 1949, F.C. Deighton M2944, IMI 37953 (K); same area, on leaves of J. multiflorum (as J. pubescens), August 1949, F.C. Deighton M3030, IMI 37954 (K). Known distribution (Fig. 7). Brazil, Costa Rica, Guyana, Puerto Rico, Suriname, Trinidad and Tobago, Venezuela (Hanlin and Tortolero 1989), Panama (new record), Sierra Leone (Deighton 1936). Notes. On leaf spots, only the lanceolate white propa- Figure 7. Sclerotium coffeicola. World distribution, countries gules were observed, which agree to the description as reported in the literature marked with green, Panama (new record) “columnar bundles of hyphae” by Hanlin and Tortolero with brown red, Sierra Leone (published record confirmed by re- (1989), but mycelia or sclerotia also described by Hanlin examination) with green color and brown red dotted arrow. and Tortolero (1989) were not found. The ITS sequence from the dried specimen from Panama had 99% similarity plant families and geographically limited to the neo- with that of the ex-type culture (CBS 115.19, GenBank tropical region, except for Africa (Sierra Leone; Farr NR_145331, Okabe and Matsumoto 2003) as well as and Rossman 2017). Re-examination of records from the with an unpublished sequence of S. coffeicola (GenBank paleotropical area revealed to represent S. rolfsii Sacc. KP176676), whereas the next similar sequences in the [= Athelia rolfsii (Curzi) C.C. Tu & Kimbr.] (Hanlin BLAST search were all from Athelia rolfsii (Curzi) C.C. and Tortolero 1989), but did not include specimens from Tu & Kimbr. with 93% similarity. In some leaf spots, Sierra Leone deposited by F.C. Deighton in K. Our study the fungus was associated with orange colored larvae of of the specimen of S. coffeicola published by Deighton gall midges (Cecidomyiidae) apparently feeding on it. (1936) only revealed sclerotia so that we were unable to The same fungus was also observed on Dioscorea alata confirm the species identification. Additional specimens, (Dioscoreaceae, material not preserved) partially grow- however, from the same locality collected by Deighton ing below and partially climbing into the infected Annona in 1949, contained the lanceolate propagules, so that we tree. On this liana, leaf spots and sporulation were sparse. could confirm the identification of the African specimens Sclerotium coffeicola is known from different host as S. coffeicola. Hanlin and Tortolero (1989) compiled Kirschner et al. | Plant parasitic fungi from Panama 99 the knowledge about the geographic distribution, hosts, to Oxalis species, but spread to economically important and heavy impact on coffee plantations, and among other plants from these weeds. The unclear taxonomy on the new information, provided descriptions of the sclero- one hand and the potential phytopathological significance tium development and ultrastructure. The ultrastructure on the other make detailed documentations of preliminar- indicated the basidiomycetous relationship, although a ily identifiable specimens particularly important as base teleomorph is not yet known. In spite of the importance for advanced studies. of coffee plantations for the economy of Panama, pub- lications about S. coffeicola on coffee leaves in Panama Acknowledgements are not known to us, whereas Mycena citricolor (Berk. & M.A. Curtis) Sacc. is a well-known threat for coffee We thank T.A. Hofmann for her collaboration concern- in Panama (Piepenbring 2006). The systematic place- ing permits and the Universidad Autónoma de Chiriquí ment of S. coffeicola in the Atheliaceae is also unclear, (UNACHI, David, Panama) for administrative support. since in spite of the available rDNA sequences, a con- J. Bernal and R. Villarreal are thanked for their help in clusive phylogenetic analysis has not been undertaken Panama and Yu-Hung Yeh for technical assistance with (Xu et al. 2010). the DNA methods. The study was financially supported by the Ministry of Science and Technology of Taiwan Discussion (NSC100-2621-B-008-001-MY3) and the SENACYT of Panama (project number APY-GC-2015-57, R. Villarreal Since knowledge about the fungal diversity in Panama and the national research program SNI). is comparatively limited, basic research providing specimens in publicly accessible collections, and detailed Authors’ Contributions descriptions and illustrations in international scientific journals as well as correctly annotated sequence data is MP and OC organized the infrastructure and its main- required for decision-making by plant protection agen- tenance for collection and study of specimens, and cies (Piepenbring et al. 2011). Because of restrictive laws identified host plants, OC arranged the collection permit, concerning export of living organisms, of lack of a public RK made the microscopy and identified the specimens, culture collection, and of limited laboratory facilities in RK and MP did the literature research and wrote the text. Panama, however, cultures could not be preserved so that DNA methods could only be applied to dried material References of the macroscopically visible propagules of Sclerotium coffeicola. For the other 2 new records from Panama, Braun U (2000) Annotated list of Cercospora spp. described by C. Annellophora phoenicis and Cercospora corniculatae, Spegazzini. Schlechtendalia 5: 57–79. Broschat TK, Elliott ML, Hodel DR (2014) Ornamental palms: biology DNA sequences would be helpful in order to address the and horticulture. Horticultural Reviews 42: 1–120. https://doi.org/ open questions of probably wrongly identified specimens 10.1002/9781118916827.ch01 of the former and the host range of the latter. Concern- Chupp C (1954) A Monograph of the Fungus Genus Cercospora. Pub- ing the doubtful records of A. phoenicis in the literature, lished by the author, Ithaca, New York, 667 pp. only the morphology of the specimen from the USA Crous PW, Braun U (2003) Mycosphaerella and its anamorphs: 1. Names published in Cercospora and Passalora. CBS Biodiversity was presented in detail by Vann and Taber (1985), so Series 1: 1–571. its identification could be corrected as Scolecostigmina Deighton FC (1936) Preliminary list of fungi and diseases of plants in palmivora, without the necessity to trace and study Sierra Leone and list of fungi collected in Sierra Leone. Bulletin the original material by ourselves. Since A. phoenicis of Miscellaneous Information (Royal Botanic Gardens, Kew) 7: belongs to quarantine organisms in some countries, e.g. 397–424. Egypt (Mohamed Ibrahim Ahmed 2010), exact species Ellis MB (1958) Clasterosporium and some allied dematiaceae-phrag- mosporae. I. Mycological Papers 70: 1–89. identification is important. Our re-examination of unpub- Ellis MB (1971) Dematiaceous Hyphomycetes. CABI Publishing, Kew, lished specimens of S. coffeicola confirmed the previous Surrey, UK, 608 pp. published record from Africa and again highlights the sci- Farr DF, Rossman AY (2017) Fungal Databases, U.S. National Fungus entific importance of curating duplicates of unpublished Collections, ARS, USDA. https://nt.ars-grin.gov/fungaldatabases. specimens (Kirschner 2016). In Cercospora, the accumu- Accessed on: 2017-11-9. lation of results of infection experiments and DNA data Groenewald JZ, Nakashima C, Nishikawa J, Shin H-D, Park J-H, Jama AN, Groenewald M, Braun U, Crous PW (2013) Species concepts showed that the concept of a host family-specific speci- in Cercospora: spotting the weeds among the roses. Studies in ficity does not work for several species so that numerous Mycology 75: 115–170. https://doi.org/10.3114/sim0012 Cercospora species have been comprised under C. apii Hanlin RT, Tortolero O (1989) Morphology of Sclerotium coffeicola, s. lat. (Crous and Braun 2003). Even in multigene stud- a tropical foliar pathogen. Canadian Journal of Botany 67: 1852– ies, several species complexes could not be resolved, but 1860. https://doi.org/10.1139/b89-234 some species appeared to be quite host specific, whereas Holubová-Jechová V (1988) Studies on hyphomycetes from Cuba VII. Seven new taxa of dematiaceous hyphomycetes. Ceska Mykologie others have broad host ranges among different plant 42 (1): 23–30. families (Groenewald et al. 2013). The species identi- Horst RK (2001) Westcott’s Plant Disease Handbook, 6th edition. Klu- fied here as C. corniculatae/C. apii may not be limited wer Academic Publishers, Dordrecht, The Netherlands, 1008 pp. 100 Check List 14 (1)

Kirschner R (2016) Alternaria bryophylli comb. nov. associated with Piepenbring M, Camarena J, Cruz D, Gomez AK, Guerrero Y, Hofmann leaf scab of Kalanchoe pinnata (Crassulaceae). Tropical Plant TA, Kirschner R, de Matas M, Perez L, Rodríguez D, Ureta J, Pathology 41: 9−14. https://doi.org/10.1007/s40858-015-0061-5 Vargas I, Williams C (2011) New records of pathogenic fungi on Kirschner R, Piepenbring M (2014) New records of three Ramichlorid- cultivated plants in Panama. Mycotaxon 115: 534−535. ium species on banana leaves in Panama and Taiwan. Mycoscience Shanthi S, Vittal BPR (2012) Fungal diversity and the pattern of fungal 55: 260−267. https://doi.org/10.1016/j.myc.2013.10.002 colonization of Anacardium occidentale leaf litter. Mycology 3 (2): Kodsueb R, McKenzie EHC, Lumyong S, Hyde KD (2008) Diversity 132−146. https://doi.org/10.1080/21501203.2011.654350 of saprobic fungi on Magnoliaceae. Fungal Diversity 30: 37−53. Thaung MM (2008) A list of hyphomycetes (and agonomycetes) in Mohamed Ibrahim Ahmed, H (2010) Seed industry in Egypt. The Afri- Burma. Australasian Mycology 27: 149−172. can Seed Trade Association. http://afsta.org/wp-content/uploads/ To-Anun C, Hidayat I, Meeboon J (2011) Genus Cercospora in Thai- documents/EGYPT%20SEED%20SECTOR%20BASELINE%20 land: Taxonomy and Phylogeny (with a dichotomous key to spe- STUDY.pdf. Accessed on: 2017-8-10. cies). Plant Pathology & Quarantine 1 (1): 11–87. Okabe I, Matsumoto N (2003) Phylogenetic relationship of Sclero- Vann SR, Taber RA (1985) Annellophora leaf spot of date palm in tium rolfsii (teleomorph Athelia rolfsii) and S. delphinii based on Texas. Plant Disease 69: 903−904. ITS sequences. Mycological Research 107: 164−168. https://doi. Xu Z, Harrington TC, Gleason ML, Batzer JC (2010) Phyloge- org/10.1017/S0953756203007160 netic placement of plant pathogenic Sclerotium species among Piepenbring M (2006) Checklist of fungi in Panama. Preliminary ver- teleomorph genera. Mycologia 102 (2): 337−346. https://doi.org/ sion. Puente Biológico 1: 1−190. 10.3852/08-189